Carbon Balance of Tall Fescue (Festuca arundinacea Schreb.): Effects of Nitrogen Fertilization and the Growing Season

The C balance of a tall fescue sward grown under different ratesof N fertilization in summer, autumn, and spring was calculatedusing models derived from measurements of shoot growth, canopygross photosynthesis, shoot respiration and of C partitioningto the roots. Under the diverse growing conditions associatedwith the seasons and the N fertilization, C utilization forabove- and below-ground biomass accumulation never exceeded39 and 14% of the canopy gross photosynthesis, respectively.Carbon losses attributed to root respiration and exudation,which were estimated by difference between canopy net photosynthesisand total growth, ranged between 3 and 30% of canopy gross photosynthesis.Seasonal differences in shoot growth could be attributed tothe amount of intercepted radiation, the radiation-use efficiencyand the C partitioning to the roots. The effect of N deficiencyon shoot growth can be attributed to its effects on canopy photosynthesis(principally resulting from changes in intercepted photosyntheticallyactive radiation) and C partitioning. In comparison with theeffect on shoot growth, the effect of the N deficiency on thecanopy gross photosynthesis per unit of light intercepted overthe regrowth cycle was limited. It is concluded that most ofthe effect of N fertilization on shoot growth is due to changesin C partitioning which result in faster leaf area developmentand greater light interception.Copyright 1994, 1999 AcademicPress Tall rescue, Festuca arundinacea Schreb., carbon balance, nitrogen, grass, fertilization     

Dynamic model of the response of a vegetative grass crop to light, temperature and nitrogen

Abstract A previously described growth model of the vegetative grass crop is extended to include a simple representation of the root system, uptake of nitrogen from a soil nitrogen pool, and response to fertilizer application. The model simulates the processes of light interception, photosynthesis, partitioning of new growth, leaf area expansion, growth and maintenance respiration, ageing of plant tissues, senescence, recycling of substrates from senescing tissues, nitrogen uptake by the plant, leaching, mineralization, and fertilizer application. A principal component of the model, nitrogen uptake, is assumed to depend positively on plant carbon substrate concentration and soil nitrogen concentration, and to be inhibited by plant nitrogen substrate concentration. The dynamic responses to different levels of soil nitrogen, of shoot and root growth, nitrogen uptake and root activity, carbon and nitrogen plant substrate concentrations, and the fraction of substrate carbon used by the shoots, are examined; realistic behaviour is observed. The model predicts nitrogen fertilizer responses of yield and plant nitrogen content, which are compared directly with experimental data; good agreement is obtained.     

Growth of Lettuce, Onion, and Red Beet. 1. Growth Analysis, Light Interception, and Radiation Use Efficiency

A field experiment was carried out to analyse the growth oflettuce, onion and red beet in terms of: (a) canopy architecture,radiation interception and absorption; (b) efficiency of conversionof absorbed radiation into biomass; and (c) dry matter partitioning.Growth analysis, total solar radiation interception, PAR interceptionand absorption by the crop canopy, ground cover, maintenancerespiration of onion bulbs and red beet storage roots were measured.Models for different leaf angle distribution and ground coverwere used to simulate light transmission by the crop canopy. The three crops are shown to have contrasting growth patternsfrom both a morphological and a physiological point of view.Lettuce showed very high light interception and growth afterthe early growth stages but, throughout the growth cycle, thisleafy crop showed the lowest radiation use efficiency due tothe respirational cost of the high leaf area. Onion showed alower early relative growth rate than lettuce and red beet.This was due partly to the low light interception per unit leafarea in the later stages of growth and partly to the low initialradiation use efficiency compared with the other two crops.On the other hand, thanks to more uniform distribution of theradiation inside the canopy, to the earlier termination of leafdevelopment and to the very low level of bulb respiration, onionshowed high radiation use efficiency and was able to producea large amount of dry matter. Red beet leaf posture and canopystructure resulted in high light interception and absorption.Its radiation use efficiency was lower than that of onion, partlyperhaps because of the more adverse distribution of the interceptedradiation fluxes within the canopy and partly because of thehigh respiration cost of a continuous dry-matter allocationto the leaves. However, this crop can accumulate a very largeamount of dry matter as leaf blade development and storage rootgrowth can both continue almost indefinitely, providing continuouslyavailable sinks. Ground cover gave a good estimate of the PAR interception onlyat low values of light interception but, in general, it underestimatedPAR interception in all three crops. Ratios between attenuationcoefficients established by considering PAR or total solar radiationand LAI or ground cover were calculated. Lettuce,Lactuca sativa L. var.crispa ; onion,Allium cepa L.; red beet; Beta vulgaris L. var.conditiva ; growth analysis; light interception and absorption; canopy architecture; ground cover; radiation use efficiency; maintenance respiration rate; dry matter distribution     

The Monomolecular and Rectangular Hyperbola as Empirical Models of the Response of Photosynthetic Rate to Photon Flux Density, with Applications to Three Veronica Species

The properties of the rectangular hyperbola and monomolecularfunctions, with respect to the photosynthesis/photon flux density(PFD) relationship, are discussed, and the shortcomings of theformer are highlighted. Both models were fitted to data acquiredfrom three closely related Veronica species of contrasting ecology.The non-linear regression algorithms give estimates, with standarderrors, of light saturated photosynthetic rate, light compensationpoint, dark respiration rate, and photochemical efficiency atlow PFD. While the rectangular hyperbola gave almost as gooda fit to the data as the monomolecular for each species, thelight saturated photosynthetic rate estimate given by the formerwas always unacceptably high in comparison with that indicatedby the obvious trend of the data. Moreover, this tendency wasaccentuated if near-saturating PFDs were removed from data sets,and there was a tendency for the fitting algorithm to becomeunstable. No such problems were encountered with the monomolecularfunction, and it is suggested that this be used whenever a simpleempirical model is required to analyze photosynthesis/PFD data. Veronica montana L, Veronica chamaedrys L, Veronica officinalis L, wood speedwell, Germander speedwell, common speedwell, empirical mathematical model, monomolecular function, rectangular hyperbola function, nonlinear regression, photosynthesis, photon flux density, light saturated photosynthetic rate, light compensation point, photochemical efficiency, dark respiration rate     

A Model of Leaf Tissue Growth, Acclimation and Senescence

A model for leaf growth, acclimation and senescence is described.The approach is mechanistic and the model, which is carbon-based,has eight state variables: substrate carbon and machinery carbon;primary, secondary and cross-linked cell wall; and two whichcorrespond to parameters of the photosynthesis light responsecurve—namely, the initial slope and light-saturated photosyntheticrate. The model is used to simulate responses to light and temperature,and predicts quantities such as final leaf area, the time atwhich the leaf becomes an exporter, senescence time, and thetwo photosynthetic parameters. The results are in good qualitativeagreement with observational responses that have been reported.The model also simulates the effects of changing the light environmentof a mature leaf: for instance, it predicts that increasingthe light level can increase leaf longevity and vice versa,although, in general, leaves grown continuously in high lighthave a shorter life than leaves grown continuously in low light. Leaf, model, growth, senescence, acclimation     

The Use of a Model to Investigate the Influence of some Environmental Factors on the Growth of Perennial Ryegrass

A modelof thegrasscrop isconstructed, inahichthe simulated valuesof crop morphology and physiology adapt automatically to changesin the environment. The model predicts photosynthesis, partitioningof assimilates and the growth or the cropin terms of leaves,stems and rmts from environmental data and the initial weightsof the crop components following defoliation. The model is usedto investigate the separate effects of light, temperature, canopystructure and maintenance respiration on the growth and denlopmcnt of the crop. It demonstrates how the crop attempts to optimizeaboveground growth, and the complexity of the interaction betweenthe physiological and environmental factors controlling growth. Lolium perenne, perennial ryegrass, photosynthesis, partition of assimilates, mathematical model     

Optimal Nitrogen Content and Photosynthesis in Cauliflower (Brassica oleracea L. botrytis). Scaling up from a Leaf to the Whole Plant

A simple model of photosynthesis is described which is dependenton leaf area, organic nitrogen content and distribution withinthe canopy as well as on the light and temperature environments.The model is parameterized using a cauliflower crop as an example.The optimized protein-N profile within the canopy is calculatedwith respect to daily growth rate. By comparison with measuredprotein-N contents, the amount of super-optimal N-uptake, i.e.the N-uptake which does not increase productivity, is assessedfor two different nitrogen and light treatments. The amountof super-optimal N accumulated in cauliflower depends on N-supplyand can exceed 80 kg N ha^-1. Copyright 2000 Annals of BotanyCompany Brassica oleracea L. botrytis, cauliflower, nitrogen, photosynthesis, respiration, model, optimization     

Dark Respiration of Winter Wheat Crops in Relation to Temperature and Simulated Photosynthesis

The dark respiration of shoots (measured between March and anthesisin mid-June) and of ears (measured between anthesis and maturityat end of July) of winter wheat crops, grown in 1982 and 1985under different nitrogen application and irrigation conditions,was determined in the field. The respiration rate of 126 averagesof four samples was measured hourly for a 12–14-h darkperiod including the night. Respiration (expressed per unitdry mass) generally declined through the season for both shootsand ears. The average rate of respiration obtained on the samenight was greater for fertilized and irrigated crops, comparedwith unfertilized and droughted crops. The relationship betweenthe measured respiration and photosynthesis, simulated usinga modified version of the model developed by Spitters (1986),was analysed. This revealed that: (a) Shoot respiration was less well correlated with photosynthesisfrom the day preceding measurement than with the average ofthe photosynthesis from the two days preceding measurement. (b) The constants relating shoot respiration to total crop photosynthesisper unit crop mass and ear respiration to total crop photosynthesisper unit ear mass had similar values. This suggests that allgrowth respiration takes place in the ears at the end of theseason. (c) Crop growth respiration consumes about 35% of assimilatebefore anthesis, and that growth respiration of the ear consumesabout 40% of assimilate at the end of the season. (d) No significant effect of treatment on the relationship betweenrespiration and photosynthesis was detected, suggesting thatthe observed effect of treatment on respiration is due entirelyto differences in photosynthesis. Triticwn aestivum var. Avalon, winter wheat, dark respiration, growth coefficient, photosynthesis model, nitrogen nutrition, irrigation     

Radiation Interception, Partitioning and Use in Grass -Clover Mixtures

Mixed swards of perennial ryegrass /white clover were grownin competition under controlled environmental conditions, attwo temperatures and with different inorganic nitrogen supplies.The swards were studied after canopy closure, from 800 to 1200°C d cumulative temperatures. Clover contents did not varysignificantly during the period. A simulation model of lightinterception was used to calculate light partitioning coefficientsand radiation use efficiencies for both components of the mixturein this controlled environment experiment. Additionally, thissame radiative transfer model was applied to the field datafrom Woledge (1988) (Annals of Applied Biology112: 175 –186)and from Woledge, Davidson and Dennis (1992) (Grass and ForageScience47: 230 –238). The measured and simulated valuesof light transmission, at different depths in the mixed canopy,were highly correlated (P<0.001) with more than 80% of thetotal variance explained. The daily average of photosyntheticallyactive radiation (PAR) interception in a natural environmentwas estimated from simulations, for the field and controlledenvironment data. Under these conditions, white clover capturedsignificantly more light per unit leaf area than perennial ryegrassat low, but not at high, nitrogen supply. In the controlled environment experiment, the radiation useefficiency of the legume was lower than that of its companiongrass. For both species, radiation use efficiency was negativelycorrelated with the mean irradiance of the leaf. The role ofa compensation between light interception and light use forstabilizing the botanical composition of dense grass –cloverswards is discussed. Light interception; radiation transfer model; growth analysis; radiation use efficiency; white clover; perennial ryegrass; Trifolium repensL.; Lolium perenneL.; grassland     

Plant growth and respiration re-visited: maintenance respiration defined - it is an emergent property of, not a separate process within, the system - and why the respiration : photosynthesis ratio is conservative

Background and Aims

Plant growth and respiration still has unresolved issues, examined here using a model. The aims of this work are to compare the model''s predictions with McCree''s observation-based respiration equation which led to the ‘growth respiration/maintenance respiration paradigm’ (GMRP) – this is required to give the model credibility; to clarify the nature of maintenance respiration (MR) using a model which does not represent MR explicitly; and to examine algebraic and numerical predictions for the respiration:photosynthesis ratio.

Methods

A two-state variable growth model is constructed, with structure and substrate, applicable on plant to ecosystem scales. Four processes are represented: photosynthesis, growth with growth respiration (GR), senescence giving a flux towards litter, and a recycling of some of this flux. There are four significant parameters: growth efficiency, rate constants for substrate utilization and structure senescence, and fraction of structure returned to the substrate pool.

Key Results

The model can simulate McCree''s data on respiration, providing an alternative interpretation to the GMRP. The model''s parameters are related to parameters used in this paradigm. MR is defined and calculated in terms of the model''s parameters in two ways: first during exponential growth at zero growth rate; and secondly at equilibrium. The approaches concur. The equilibrium respiration:photosynthesis ratio has the value of 0·4, depending only on growth efficiency and recycling fraction.

Conclusions

McCree''s equation is an approximation that the model can describe; it is mistaken to interpret his second coefficient as a maintenance requirement. An MR rate is defined and extracted algebraically from the model. MR as a specific process is not required and may be replaced with an approach from which an MR rate emerges. The model suggests that the respiration:photosynthesis ratio is conservative because it depends on two parameters only whose values are likely to be similar across ecosystems.     

Growth Response of a Chrysanthemum Crop to the Environment. III, Effects of Radiation and Temperature on Dry Matter Partitioning and Photosynthesis

Vegetative crops of chrysanthemum were grown for 5 or 6 weekperiods in daylit assimilation chambers. Crop responses to differentradiation levels and temperatures were analysed into effectson dry matter partitioning, specific leaf area, leaf photosynthesisand canopy light interception. The percentage of newly formed dry matter partitioned to theleaves was almost constant, although with increasing radiationor decreasing temperature, a greater percentage of dry matterwas partitioned to stem tissue at the expense of root tissue.There was a positive correlation between the percentage of drymatter in shoot material and the overall carbon: dry matterratio. Canopy photosynthesis was analysed assuming identical behaviourfor all leaves in the crop. Leaf photochemical efficiency wasonly slightly affected by crop environment. The rate of grossphotosynthesis per unit leaf area at light saturation, P_A (max),increased with increasing radiation integral, but the same parameterexpressed per unit leaf dry matter, P_w (max) was almost unaffectedby growth radiation. In contrast, P_A (max) was hardly affectedby temperature but P_w (max) increased with increasing growthtemperature. This was because specific leaf area decreased withdecreasing temperature and increased with decreasing radiation.There was a positive correlation between canopy respirationintegral and photosynthesis integral, and despite a four-foldchange in crop mass during the experiments, the maintenancecomponent of canopy respiration remained small and constant. Canopy extinction coefficient showed no consistent variationwith radiation integral but was negatively correlated with temperature.This decrease in the efficiency of the canopy at interceptingradiation exactly cancelled the increase in specific carbonassimilation rate that occurred with increasing growth temperature,giving a growth rate depending solely on the incident lightlevel. Chrysanthemum, dry matter partitioning, photosynthesis, specific leaf area     

An Evaluation of the Effects of Ethylene on Carbon Assimilation in Lycopersicon esculentum Mill

Leaf and whole plant gas exchange rates of Lycopersicon esculentumMill, were studied during several days of continuous exposureto ethylene. Steady-state photosynthesis and transpiration ratesof control and ethylene-treated individual leaves were equivalent.However, the photosynthesis and transpiration rates of treatedleaves required at least five times longer to reach 50% of thesteady-state rate. This induction lag was attributed to ethylene—inducedleaf epinasty and temporary acclimation to lower incident lightlevels immediately prior to measurement of gas exchange. Thewhole plant net carbon exchange rate (NCER) of a representativetreated plant was also reduced by 51% after 24 h exposure toethylene relative to both its pre-treatment rate and that ofthe control. Ethylene exposure reduced the growth rate of thetreated plant by 50% when expressed as carbon (C) gain. Theinhibition of NCER and growth rate associated with epinastywas completely reversed when the epinastic leaves were returnedto their original positions and light interception was re-established.The results demonstrate that the inhibition of whole plant CO₂assimilation is indirect and due to reduced light interceptionby epinastic leaves. Morphological changes caused by environmentalethylene are thus shown to reduce plant C accumulation withoutinhibiting leaf photosynthesis processes per se. Key words: Ethylene, carbon assimilation, growth     

Tradeoffs Between Irradiance Capture and Avoidance in Semi-arid Environments Assessed with a Crown Architecture Model

Plants in arid environments cope with stress from excessiveirradiance by physiological photoprotection of the photosyntheticapparatus and by structurally reducing the leaf area exposedto the sun (structural photoprotection). We assessed the ecologicalrelevance of structural photoprotection in two plants of contrastingarchitecture co-occurring in a semi-arid environment, usingthe three-dimensional canopy model YPLANT. We compared the roleof crown geometry in avoiding excessive radiation, analysedthe costs of structural photoprotection in terms of reductionof potential carbon gain, and compared these costs with thosedue to seasonal constraints of photosynthesis and tissue ageing.The results of the model simulations indicated that canopy architectureofStipa tenacissima(a tussock grass) andRetama sphaerocarpa(aleafless leguminous shrub) minimized the risk of overheatingand photo-oxidative destruction of the photosynthetic apparatuswith steeply oriented foliage and moderate self-shading. Butthis structural photoprotection imposed an increased cost interms of potential carbon gain. Diurnal and seasonal patternsof light interception by the crown of these plants translatedinto a simulated potential carbon gain only half that of anequivalent, horizontal photosynthetic surface. This reductionin potential carbon gain, due to irradiance avoidance, was similarto that imposed by water shortage.S. tenacissima,which ceasesphotosynthetic activity during periods of drought, exhibitedmore structural avoidance of irradiance thanR. sphaerocarpa,whichremains active throughout the year. This illustrates the influenceof the capacity of plants to utilize light for carbon fixationon the trade-offs between irradiance capture and avoidance.Structural avoidance of excessive radiation efficiently preventsthe risk of damage by intense irradiance, has no special maintenancecosts, and is biomechanically cheaper than enhanced light harvestingby a horizontal canopy, which points to structural photoprotectionas a very effective strategy to cope with high irradiance stressin poor and adverse habitats.Copyright 1999 Annals of BotanyCompany Canopy model, functional architecture, irradiance avoidance, light harvesting, photoinhibition, photoprotection, photosynthesis model, plant architecture,Retama sphaerocarpa,semi-arid environment,Stipa tenacissima,structural photoprotection.     

Canopy Photosynthesis and Crop Growth Rate of Eight Temperate Forage Grasses

The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q₁₀ = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.     

Regrowth by Swards of Subterranean Clover after Defoliation. 2. Carbon Exchange in Shoot, Root and Nodule

The carbon economy of subterranean clover swards subjected tothree defoliation treatments (removal of 30, 70 and 80% of shootdry weight) was compared with that of uncut swards. Carbon dioxideexchange in shoots and roots was measured independently 0, 4,8 and 12 d after defoliation. The respiration linked to nitrogenaseactivity was estimated by comparing root respiration measuredin an atmosphere containing 3% oxygen with the respiration in21% oxygen. Net photosynthesis fell by up to 100% immediately after defoliation.There was a decline of over 60percnt; in root respiration bythe end of the first light period, composed of a rapid declineof 70% in nitrogenase-linked respiration in all treatments anda slower decline of nearly 40% in root plus nodule growth andmaintenance respiration in the more severe treatments. Recoveryof net photosynthesis to rates achieved by uncut swards occurredover 4 d in the 30% cut treatment and at least 12 d in moresevere treatments. Whilst recovery of photosynthesis was theprinciple determinant of recovery of net positive carbon balance,the early reduction in respiration facilitated this outcome.After the immediate decline in nitrogenase-linked respiration,recovery in this component of respiration appeared to be linkedwith the recovery in net photosynthesis (approximately 10% ofnet photosynthesis). Carbon budgets revealed priorities in allocation towards leafin the first 5 d and later also towards root growth in severelydefoliated swards. Root respiration comprised a large respiratorycost (up to 75% of net photosynthesis) during early regrowth. Carbon budget, defoliation, N₂ fixation, photosynthesis, regrowth, respiration, subterranean clover, Trifolium subterraneum L     

LIGNUM: A Tree Model Based on Simple Structural Units

The model LIGNUM treats a tree as a collection of a large numberof simple units which correspond to the organs of the tree.The model describes the three dimensional structure of the treecrown and defines the growth in terms of the metabolism takingplace in these units. The activities of physiological processescan be explicitly related to the tree structures in which theyare taking place. The time step is 1 year. The crown of the model tree consists of tree segments, branchingpoints and buds. Each pair of tree segments is separated bya branching point. The buds produce new tree segments, branchingpoints and buds. The tree segments contain wood, bark and foliage.A model tree consisting of simple elements translates convenientlyto a list structure: the computer program implementing LIGNUMtreats the tree as a collection of lists. The annual growth of the tree is driven by available photosyntheticproducts after respiration losses are accounted for. The photosyntheticrate of foliage depends on the amount of light. The amount ofphotosynthates allocated to the growth of new tree segmentsis controlled by the light conditions and the amount of foliageon the mother tree segment. In principle, the biomass relationshipsof the tree parts follow the pipe model hypothesis. The orientationof new tree segments results from the application of constantbranching angles. LIGNUM has been parametrized for young Scotspine (Pinus sylvestrisL.) trees. However, the model is generic;with a change of parameter values and minor modifications itcan be applied to other species as well. Growth model; object-oriented modelling; tree architecture; branching structure; Pinus sylvestrisL.; developmental morphology and physiology; photosynthesis; respiration     

Effect of Temperature and Nitrogen Supply on the Growth of Perennial Ryegrass and White Clover. 1. Carbon and Nitrogen Economies of Mixed Swards at Low Temperature

Simulated mixed swards of perennial ryegrass (Lolium perenneL. cv. S23) and white clover (Trifolium repens L. cv. S100)were grown from seed under a constant 10°C day/8°C nighttemperature regime and their growth, and carbon and nitrogeneconomies examined. The swards received a nutrient solution,every second day, which contained either high (220 µgg^–1) or low (40 µg g^–1) nitrate N. The High-N swards had rates of canopy photosynthesis and drymatter production (over the linear phase of growth) similarto those previously shown by mixed swards at high temperature.The Low-N swards grew more slowly; canopy photosynthesis, ata given LAI, was similar to that at High-N but lower LAI's weresustained. Clover increased its contribution to total carbonuptake and total dry weight throughout the period in the Low-Ntreatment and, despite the fact that grass took up most of theavailable nitrate, clover maintained a consistently higher Ncontent by virtue of N₂-fixation. At High-N, grass dominated throughout the measurement period.Earlier, when plants grew as spaced individuals, clover grewless well than grass, but once the canopy was closed it hada similar relative growth rate and thus maintained a steadyproportion of total sward dry weight. It is proposed that earlyin the development of the crop, leaf area production is thelimiting factor for growth, and that in this respect cloveris adversely affected by low temperature relative to grass.Later, as the LAI of the crop builds up, and the canopy becomesfully light intercepting, net canopy photosynthesis plays amore dominant role and here the higher photosynthetic rate perunit leaf area of the clover is crucial. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, low temperature, nitrogen, photosynthesis     

Photosynthesis by White Clover Leaves in Mixed Clover/Ryegrass Swards

Measurements of rates of net photosynthesis were made on singleBlanca white clover leaves on plants taken from a field-grown,mixed clover/perennial ryegrass sward during two regrowth periods. Net photosynthesis fell by 20 per cent in the first measurementperiod as leaf area index increased and the grass componentof the crop flowered, but did not change significantly in thesecond measurement period during which the grass remained vegetative. Leaves which had been artificially protected from shading inthe sward did not have significantly different photosyntheticcapacities from leaves in the undisturbed sward, even in thefirst measurement period. As leaf area index and sward height increased, successive cloverpetioles were longer, keeping the newly expanded leaves nearthe top of the sward where they received full light. It is suggestedthat it is this which allows successive clover leaves, unlikethose of vegetative grasses, to attain a high photosyntheticcapacity throughout a growth period. Trifolium repens, Lolium perenne, Photosynthetic capacity, shading, growth     

Effect of Nitrogen Supply on the Grass and Clover Components of Simulated Mixed Swards Grown under Favourable Environmental Conditions I. Carbon Assimilation and Utilization

Simulated mixed swards of Perennial Ryegrass (Lolium perenneL.) cv. S23 and White clover (Trifolium repens L.) cv. S100were grown from seed under a constant 20 °C day/15 °Cnight temperature regime and their growth and carbon economyexamined. The swards received a nutrient solution daily, whichcontained either High (220 mg l^–1) or Low (10 mg l^–1)nitrate N. Rates of canopy photosynthesis and respiration, and final drymatter yields were similar in the two treatments although theproportions of grass and clover differed greatly. The Low-Nswards were made up largely of clover. The grass plants in theseswards had high root: shoot ratios and low relative photosyntheticrates – both signs of N deficiency – and were clearlyunable to compete with the vigorously growing Low-N clover plants.These had higher relative growth rates and dry matter yieldsthan their High-N counterparts. In the High-N swards clovercontributed around 50 per cent to the sward dry weight throughoutthe measurement period despite having a smaller proportion ofits dry weight in photosynthetic tissue (laminae) than grassover much of it. The latter was compensated for, initially bya higher specific leaf area than grass, and later by a higherphotosynthetic rate per unit leaf weight. The results are discussedin relation to observed declines in the clover content of swardsafter the addition of nitrogen fertilizer in the field. Trifolium repens, white clover, Lolium perenne, perennial ryegrass, nitrogen, photosynthesis, carbon balance     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.